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Contents JOURNAL OF THE NETHERLANDS MALACOLOGICAL SOCIETY BASTERA VOLUME 82 (4-6) | 30 DECEMBER 2018 VOLUME 82 (4-6) | 30 DECEMBER 2018

Vermeij, G.J. & Snyder, M.A. Proposed genus-level classification of large of Fusininae ( , Fasciolariidae) ...... 57 Janssen, A.W., Burridge, A.K., Mekkes, L. & Peijnenburg, K.T.C.A. History of the pyramidata -complex (, Gastropoda, , Cliidae) in the Atlantic Ocean , restoring its original concept ...... 83 Jagt, J.W.M. Latest Cretaceous Pulvinites (Bivalvia, Pterioidea, Pulvinitidae) , new to the Maastrichtian type area, The Netherlands ...... 103 Janssen, A.W. Notes on the systematics, morphology and biostratigraphy of holoplanktic Mollusca, 25 . Once more: the correct name for the type species of the genus Creseis Type specimens of Rang, 1828 (, Euthecosomata, Creseidae) ...... 110 Fusininae (p.57) Basteria_cover_82(4-6):Opmaak 1 12/2/2018 4:44 PM Page 2

Editorial board

Prof. Dr E. Gittenberger (editor-in-chief), Dr B. Kokshoorn (layout editor), G.C. Kronenberg (editor).

Associated editors

Prof. Dr G. van der Velde (freshwater molluscs), Prof. Dr G.J. Vermeij (fossil and recent marine molluscs), Dr F.P. Wesselingh (fossil molluscs).

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History of the -complex (Mollusca, Gastropoda, Euthecosomata, Cliidae) in the Atlantic Ocean , restoring its original concept

Arie W. Janssen Naturalis Biodiversity Center, Marine Biodiversity, P.O. Box 9517, NL-2300 RA Leiden, The Netherlands; [email protected] [corresponding author]

Alice K. Burridge Institute for Biodiversity and Ecosystem Dynamics (IBED), University of Amsterdam, P.O. Box 94248, NL-1090 GE Amsterdam, The Netherlands ; [email protected]

Lisette Mekkes Naturalis Biodiversity Center, Marine Biodiversity, P.O. Box 9517, NL-2300 RA Leiden, The Netherlands . Institute for Biodiversity and Ecosystem Dynamics (IBED), University of Amsterdam, P.O. Box 94248, NL-1090 GE Amsterdam, The Netherlands; [email protected]

Katja T.C.A. Peijnenburg 83 Naturalis Biodiversity Center, Marine Biodiversity, P.O. Box 9517, NL-2300 RA Leiden, The Netherlands. Institute for Biodiversity and Ecosystem Dynamics (IBED), University of Amsterdam, P.O. Box 94248, NL-1090 GE Amsterdam, The Netherlands; [email protected]

‘My heart is down, my head is turning around ’ (Harry Belafonte, Jamaica Farewell, 1956)

An extensive overview of the historical use of the genus currently indicated as forma or subspecies lanceolata Clio and its type species Clio pyramidata Linné, 1767 (de - Lesueur, 1813 . Based on combined morphological and scribed from the Caribbean: Jamaica) during the last genetic analyses of specimens from the Atlantic Ocean two and a half centuries demonstrates the current in - and the Caribbean we found that there are intermedi - correct use of the species name and the junior syn - ates between the two shell forms and that they have onymy of the taxon Hyalea lanceolata Lesueur, 1813 different, but overlapping distributions. Moreover, (described from the Mediterranean). The latter taxon there was no molecular evidence to separate the two was incorrectly considered to be a shell-morphologi - forms as distinct species. Hence, we consider the name cally different form of C. pyramidata . A neotype is desig - C. pyramidata appropriate for the different shell forms nated for C. pyramidata from the original type locality to within this taxon. For the form with straight carinae the stabilise nomenclature. We examined the relationship name angusta Boas, 1886 , is available, but may be ap - of two shell forms of that species, viz. the form with plied at subspecific rank only. A lectotype for angusta is straight carinae or shell sides (occurring in the northern selected, and a type species is designated for Flabellu - Atlantic Ocean and so far considered to represent typi - lum Bellardi, 1873. cal C. pyramidata ) and the (neo)typical form with strongly concave carinae (occurring in the (sub)tropical Key words: systematics, nomenclature, distribution, type local - Atlantic, inclusive of the Caribbean and Mediterranean) ity, neotype, lectotype, barcoding, Jamaica.

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Introduction 1774 : 195) indeed does not at all represent a species with a compressed shell (which is Linné’s description The euthecosomatous pteropod Clio pyramidata Linné, of 1C. caudata ), but is a completely different pteropod 1767 , is one of the more abundant holoplanktic mol - species, without a shell and belonging to the Gymno - lusc species with a very wide distribution in almost all somata, but very much agreeing with Linné’s diagno - oceans and seas (e.g., van der Spoel, 1967) . The species sis of the genus Clio. Linné’s diagnosis and the is, generally speaking, easily recognisable, although reference to Martens’ paper led to much confusion as its wide distribution in cold as well as warm waters several later authors decided to include the same or has led to the introduction of a number of forms similar shell-less pteropod species in the genus Clio. and/or subspecies , or even species, based on shell morphology . In the Atlantic two different shell forms are present, distinguished by the shape of the carinae (shell sides) in dorsal and ventral views: straight in the North Atlantic, more concave in the (sub) tropical At - lantic, the Caribbean and Mediterranean. Over the course of time , erroneous interpretations have changed the original species concept , resulting in the current incorrect application of the name. Here we provide a historical survey of the taxon and, in the ab - sence of syntype material, restore its original defini - tion by designation of a neotype from its type locality Fig. 1. Martens‘ (1675: 128, pl. P figs f) ‘See Gots-Pferd’ = Clione Jamaica. Furthermore , we examined the different limacina (Phipps, 1774). Size not indicated (maximum body forms occurring in the northern and central Atlantic length up to 40 mm). Ocean by means of combined analyses of shell form and DNA sequencing . 3 Martens ’ species was, shortly after Linné, formally de - Abbreviations: NHMD = Natural History Museum of scribed as Clio limacina Phipps, 1774, and also as Clione 84 Denmark, Copenhagen; RGM = Rijksmuseum van Ge - borealis Pallas, 1774, of which the former name has pri - ologie en Mineralogie, collection now in Naturalis Bio - ority (MolluscaBase, 2018a ). Müller (1776 : 226 , nr diversity Center, Leiden, The Netherlands. 2742 ), however, apparently not aware of Phipps’ and Pallas’ papers, used Linné’s name Clio retusa for this Historical notes on the genus Clio species. He added to the confusion by including the short Latin description of the shell given by Browne Linné (1767 : 1094), when formally introducing the and repeated by Linné ( ‘vagina triquetra, ore horizon - genus Clio , a name borrowed from Browne (1756 : 386), tali ’), but he also added the reference to Martens diagnosed the genus as ‘ Corpus natans, oblongum; Alis (1675) and the Greenlandic vernacular names of the duabus, membranaceis, oppositis ’, which describes the species described by Martens ( ‘Ataurfak, Augurfak ’) . pelagic way of life, the shape of the body and the pres - Also Fabricius (1780 : 334) used the name Clio retusa for ence of two opposite , membranaceous parapodia , but this gymnosome species and gave an extensive de - does not mention the presence of a shell. Linné, how - scription, but realising that this is a shell-less species, ever, included only the three -shelled species already he left ‘vagina triquetra’ out of his diagnosis. Fabricius described but unnamed by Browne (1756: 386), for referred to Pallas’ 1774 description, but cited for that which he introduced the names Clio caudata, C. pyrami - reference the incorrect name Clione papilionacea , not data and C. retusa . For each of these a shell (Latin: used by Pallas. ‘vagina ’) is described, following Browne’s earlier de - Gmelin (1791) also included Clio limacina Phipps scriptions. For C. caudata , diagnosed as ‘C . vagina com - and Clione borealis Pallas in the genus Clio , and trans - pressa caudata ’, a taxon currently considered to be a ferred the reference to Martens from Clio caudata to synonym of C. pyramidata , Linné not only referred to Clio limacina , while retaining the description of the C. Browne’s (1756) description, but also included a sug - caudata shell as given by Browne and Linné, adding gestion (‘ Conf. ’ = compare) to Martens (1675 : 128, pl. P references to Müller and Fabricius. This mixing up of fig. f), adding the remark ‘qui nullam vaginam in sua de - completely different species has caused even further lineavit ’, which seems to mean ‘ who did not mention a confusion in later literature . shell in his text ’. Bruguière (1792 : xi) included the genus Clio in his Martens’ illustration (copied here as Fig . 1) of his ‘Tableau systématique des vers , Ordre III – Les vers mol - ‘See Gots-Pferd ’ ( translated as ‘sea may-flye’ in Phipps, lusques , section II - Avec tentacules’ , containing exclusi -

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vely shell-less species. On pages 505-508 he discussed 20, etc. ) indeed applied the genus Clio for the shelled the genus Clio , considering the genus Clione Pallas, species, but other authors (e.g., Gegenbaur, 1855 : 210; 1774 , to be a synonym of Clio Linné, 1767, and accept - Conti, 1864 : 26; Seguenza, 1867 : 9; Weinkauff, 1868 : ing only Clio borealis Pallas, 1774 , and C. australis 425-426; Bellardi, 1873 : 30; Fol, 1873 : 177; Fischer, Bruguière, 1792, both belonging to the Gymnoso - 1883 : 435; Boas, 1886 : 54; Friele & Grieg, 1901 : 122 ; mata, and currently part of the genus Clione (fide van Collins, 1934 : 200) retained the name Cleodora as der Spoel, 1976: 110, 115). Bruguière excluded the valid. shelled species of Browne and Linné from the genus Bonnevie (191 3: 2 0) introduced the subgenus Eu - Clio , as according to him they belonged to another clio in Clio for three species, among which also Clio group, namely his ‘Ordre V – les vers testacés ’ and an - pyramidata. This makes Euclio an objective junior syn - nounced their future description ‘ dans la suite au mot onym of Clio . fissurelle [ ‘in a next part under the word fissurelle’] ’. Thiele’s (1931 : 401) ‘Handbuch der systematischen Cuvier (1802 : 242-244 ) described this ‘chaos ’ and Weichtierkunde ’ listed Clio as a valid genus in the stated that the species described as Clione borealis by , with Cleodora , Balantium Children, 1823, Pallas had become the most commonly known and and Euclio as synonyms, and Clio pyramidata as type had to be considered the type species of the genus Clio . species. Tesch (1946: 13-14) , however, retained Euclio , This statement, however, cannot be taken as valid, as and although he was well aware of ‘the rules of prior - the taxon C. borealis Pallas was not included in Linné’s ity’ (Tesch, 1946: 14) also wrote ‘seems to me prefer - description of Clio. Such taxonomic ruling, however, able to Pelseneer’s Clio (s. str. )’. did not yet exist in the early nineteenth century . Hubendick (1951 : 3) introduced the genus Proclio Probably it was Cuvier’s unchallenged authority for the species P. subteres Hubendick, 1951, that was that also led Péron & Lesueur (1810 : 65 ) to arrange subsequently synonymised with Clio pyramidata f. the genus Clio among the group of pteropods without antarctica Dall, 1908 , by van der Spoel (1967 : 72 ; see a shell, which at the same time necessitated them to also van der Spoel, 1962), which made Proclio a sub - introduce a new genus , Cleodora Péron & Lesueur, jective junior synonym of Clio. However, Clio pyrami - 1810 , for the shelled species described by Browne and data f. antarctica was found to be genetically distinct Linné. They included a single species in the new from C. pyramidata by Burridge et al. (2017 , fig. S1 ), genus, indicated with the vernacular name ‘Cléodore which makes the name Proclio available again. 85 Pyramidale’ (their pl. 2 fig . 14) (see next chapter ). The Zilch (1959 : 51 fig . 169) also placed Cleodora and illustration is rather poor and schematic, but recog - Euclio in synonymy with Clio , together with Balantium nisable as Clio pyramidata as interpreted until that and some taxa introduced by Bellardi (1873) for fossil time, and showing a triangular shell form with com - species (Poculina, Flabellulum ). That author, however, pletely straight carinae . retained the genus Proclio as valid. His illustration is After 1810, the name Clio was predominantly ap - copied from Sowerby (in Reeve, 1878, Pteropoda pl. 4 plied to the shell-less gymnosome species, and fig. 25a-b) and represents a specimen of Clio pyrami - Cleodora was commonly used for the shell-bearing data with concave carinae . As type species of Clio species. Herrmannsen (184 7: 247-250) gave many ref - Zilch erroneously mentioned Clio ‘pyramidalis Lin - erences for the use of the names Clio and Cleodora . naeus ‘, obviously a lapsus calami for pyramidata . The Bruguière’s 1792 promise to describe Browne’s and same error occurs in the caption of his fig. 169. Also Linné’s shelled species later was only realised in 1830, Bowdich (1822) had described a ‘Cleodora pyramidalis ’. in the second volume of the Histoire naturelle des vers , Confusingly , the name Clio pyramidalis had also and then under the authorship of Deshayes. In the validly been introduced for a gymnosome pteropod meantime the genus Cleodora was introduced and one by Quoy & Gaimard (1832), see next chapter . might have expected that the shelled species were In van der Spoel (1967: 65-66 ) the names Cleodora, discussed there. However, only Lesueur’s 1813 Balantium, Euclio and also Proclio are considered syn - species Hyalea lanceolata was included there with the onyms of Clio. Also an extensive synonymy is given name Cleodora lanceolata , not a word on Browne’s and for this genus This author accepted nine species in Linné’s species. In that publication the genus Clio , is the genus Clio and additionally seven formae . still used for the gymnosome species only. The application of subgenera in the genus Clio At last Gray (1847 : 203) synonymised Cleodora was discussed by Janssen (2000 : 44), who observed Péron & Lesueur, 1810 , with Clio Linné, 1767 , and that in the various recent as well as fossil species not designated Clio pyramidata as the type species of Clio . only considerable differences in adult shell morphol - Thereafter several authors (e.g., Adams & Adams, ogy occur, but also in the details of the protoconchs. 1853 : 52; Philippi, 1853 : 291; Jeffreys , 1869: 118; Pelse - He suggested the use of Clio s. str . and Balantium as neer, 1888 : 42; Norman, 1890 : 62 ; Meisenheimer, 1905 : subgenera, but noted that other Recent species do not

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conform to the morphology of these two. Later From the illustration it is clear that the artist had seen (Janssen, 2005) the subgenus Bellardiclio was intro - a live , as the parapodia are drawn in life posi - duced for C. cuspidata Bosc , 1801, and a related fossil tion. Several later authors, however, criticised the rep - species. Assigning most fossil cliids to subgenera, resentation of the animal’s head. The shell is drawn as however, is problematic as in many of the species de - a regular triangle, with completely straight carinae scribed from the fossil record the protoconchs are still and, agreeing with Browne’s description, an obliquely unknown. Currently (e.g. , in WoRMS) such subgenera positioned triangular aperture. However, a form of C. in Clio are not accepted, which may be changed once pyramidata with such straight carinae does not occur in sufficient molecular data have become available, the type area (here all specimens would have concave which may lead to accepting these taxa at full genus sidelines ). We have arrived at the inevitable conclu - level. The genus Clio was one of the genera that was sion that it is especially this picture, together with sev - not found to be monophyletic in molecular phyloge - eral follow-ups of later authors, that have initiated an netic analyses of Corse et al. (2013) and Burridge et al. erroneous interpretation of the species Clio pyramidata (2017). that has generally been accepted for more than 250 Only in 2006 the International Commission on Zo - years, as we will explain, and for which we introduce ological Nomenclature ruled that the name ‘ Clio Lin - a solution , below . naeus, 1767’ is placed on the Official List of Generic A second early illustration was published , as an Names in Zoology, with its type species ‘ Clio pyrami - unnamed ‘insect’ , by La Martinière (1 787 a: 207 , pl. 11 data Linnaeus, 1767’ , by subsequent designation of figs 1-3 ), observed during the unhappy voyage of De Gray (1847). Currently eight recent Clio species are ac - la Pérouse , near Notka (Vancouver Island, Canada ; cepted (MolluscaBase 2018b) . Subgenera are not used. Fig . 3). Also these drawings are apparently based on The various earlier distinguished formae have no offi - the observation of a live animal, and although show - cial status, as ruled by ICZN (1999). ing several differences compared to Browne’s draw - ing, it unambiguously represent the same species . The species Clio pyramidata in the eighteenth and nineteenth century

86 The species Clio pyramidata , together with C. caudata and C. retusa , were formally introduced by Linné (1767 : 1094), who obviously had no specimens before him but based the names on the descriptions of Browne (1756 : 386), who had only illustrated C. pyra - midata in his pl. 43 fig . 1 ( reproduced herein as Fig . 2). Syntypes are not available either . The short descrip - tions of Browne, repeated by Linné, leave no doubt that all three names represent shelled species. Fig. 3. Unnamed ‘insect’, after La Martinière, 1787a, from ‘près de Notka’ (Vancouver Island, British Columbia, Canada). Size not indicated.

Gmelin (1791 : 3148) followed Linné (1767) in includ - ing the same three species in the genus Clio that were adopted from Browne (1756) , but adding references for Clio retusa to Müller (1776) and Fabricius (1780) , which both, however, concerned unshelled gymno - some species , and giving a rather long Latin descrip - tion for that species, at least partly based on Fabricius ’ even longer description. Apart from the three Linnean species , Gmelin furthermore included Clio borealis Pal - las , C. limacina Phipps, which again are unshelled Fig. 2. Browne’s (1756) illustration of Clio pyramidata , from Ja - gymnosomes , and C. helicina Phipps. For the latter maica (enlarged, the size was indicated as ‘length seldom ex - Gmelin referred also to Martens (1675 : 129, pl. Q fig. ceeds half an inch’. e), where a so-called ‘ Schnecken Rotzfisch ’ (‘sea slime- fish ’ in an English translation, White, 1855) was de -

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scribed and illustrated. These three drawings (here Ten years after its first publication , the same paper of La reproduced in Fig. 4), however, evidently represent a Martinière (1787a) was republished posthumously and shelled, sinistrally coiled pteropod in apical view, almost unchanged in volume 4 of the ‘ Voyage de la with the parapodia indicated. This species is cur - Pérouse ’ (La Martinière, 1797 : 61, pl. 20 figs 1-3 ), but with rently known as Limacina helicina (Phipps, 1774) and redrawn, even more unrealistic illustrations (Fig . 6). has nothing to do with Clio . Martens’ illustrations may be the earliest recognisably published euthecoso - matous pteropod.

Fig. 4. Martens’ (1675, pl. Q fig. e) illustrations of ‘ Schnecken Ro4 - Fig. 6. Unnamed ‘insect’, after La Martinière, 1797, from Nootka fisch ’ = Limacina helicina (Phipps, 1774). (Vancouver Island, British Columbia, Canada).

Gmelin (1791: 3148) also included a reference to La Bosc (180 1: 241, pl. 9 figs 1-3; Fig . 7 herein) recognised Martinière (1787b : 366 , pl. 2 figs 15-16 ) for Clio li - La Martinière ’s (1787a ) illustrations as the species ear - macina , where again an unnamed ‘insect’ was de - lier figured by Browne and described as Clio pyrami - scribed. La Martinière’s illustration (Fig . 5 herein ), data by Linné. Bosc included it in the genus Hyalæa however, does represent not even a pteropod, but a Lamarck (1799: 89) , a monotypic genus introduced pelagic nudibranch , Glaucus atlanticus , already de - for Anomia tridentata Forsskål in Niebuhr, 1775, which 87 scribed (but not illustrated) by Forster (1777: 49) and currently is included in the genus Cavolinia . He gave so beautifully pictured on the cover of Lalli & new figures that evidently are based on the earlier Gilmer’s (1989) book. Gmelin’s decision to incorpo - published drawings of La Martinière, but have again rate it in the genus Clio is explained by a footnote in been changed and further deviate from reality. Sev - La Martinière (1787b : 366) saying ‘ Il paroit se rap - eral later authors also preferred to use the genus procher du genre des clios ’ [ it seems to be close to the name Hyalæa , but see the extensive description and genus clio ]. Although the genus Glaucus was known discussion in de Blainville (1821). to Gmelin (1791 : 3025), he did not recognise La Mar - To the description of their new genus Cleodora , tinière’s figures to belong to it. Péron & Lesueur, 1810, pl. 2 fig . 14, reproduced herein as Fig . 8, added an illustration of the only in - cluded species of that genus, indicated as ‘Cléodore Pyramidale’, a schematised drawing very much re - sembling the original figure of Browne (1756).

Fig. 5. Unnamed ‘insect’ in La Martinière (1787b, pl. 2 figs 15-16) from Baschi (?) Islands = Glaucus atlanticus Forster, 1777 (Nudi - Fig. 7. Hyalæa pyramidata , after Bosc (1801), from ‘les mers branchia). d’Amérique’.

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tilo-gelatinous; like a reversed truncated pyramid ’, and gave its distribution as ‘Seas of warm climates’. His illustration of the ‘envelope’ of this species is rather schematic, showing a shell with completely straight sidelines, that seems to be based on the early drawing in Browne (1756) . Bowdich’s illustration is copied here in Fig . 10.

Fig. 8. Péron & Lesueur’s (1810) illustration of ‘Cléodore pyrami - dale’.

From the Mediterranean , off Nice , Lesueur (1813 : 284, pl. 5 fig. 3) introduced a new species, described as Fig. 10. The ‘envelope’ of Cleodora ‘pyramidalis’ , after Bowdich Hyalea lanceolata . Curiously, the author did not include (1822, pl. 5 fig. 2). Enlarged, size and locality not mentioned. his new species in the genus Cleodora , introduced few years earlier (1810) by Péron and himself. The draw - ings (copied herein as Fig . 9 ) show a Clio resembling A new taxon was introduced by de Blainville ( 1824: the earlier illustrations of C. pyramidata , but differing 272; 1825: 481; 1827, pl. 46bis figs 1, 1a-b) with the by the strongly diverging , concave carinae of the shell, name ‘Cléodore de Browne, Cleodora Brownii ’, obvi - 88 ending in short lateral spines at the aperture . ously based on Browne’s (1756) illustration. He gave a short description and referred to Péron & Lesueur (1810, pl. 3 fig. 14) (error for pl. 2), which is ‘Cléodore pyramidale’ - Clio pyramidalis . He described the Cleodora shell as ‘ une sorte d'étui gélatineux à ouverture antérieure fort grande, et non échancrée latéralement [sort of a gelatinous case with a very large anterior open - ing, without lateral slits]. His drawings are repro - duced here as Fig . 11. Van der Spoel (1967 : 68) synonymised Cleodora Fig. 9. Hyalea lanceolata Lesueur, 1813. After Lesueur (1813: 284, brownii with Clio pyramidata , but this seems to be a bit pl. 5 fig. 3A-C). doubtful because of the rather strong lengthwise cur - vature of the shell as seen in de Blainville’s fig . 1 .a (Figure . 11 herein). In Clio pyramidata the shell is virtu - Bowdich (1822 : 23, 56, pl. 5 fig. 2) attributed the genus ally straight. Cleodora to Péron (not Péron & Lesueur) and included From the voyage of the ‘Astrolabe’ Quoy & Gaimard it , together with some other genera , in a new division (1832 : 386, pl. 27 figs 7-13) gave a description and illus - ’Vaginaceae’ of his ‘Univalve shells’. The division trations of Cleodora pyramidata from the southern At - name is based on the fossil pteropod genus Vaginella lantic Ocean , referring to Hyalea lanceolata Lesueur as a Daudin , 1800, that Bowdich, however, renamed synonym. In their description they give a number of Vaginellites , as was frequently done for fossils. He de - anatomical soft part details . Their illustration repre - scribed Cleodora ‘pyramidalis’ (lapsus calami) 1 as ‘car - sents a shell with concave sidelines . Rather confusingly, however, they also (Quoy & Gaimard, 1832: 371, pl. 27 1 This author appears to have been rather shabby in his namings, fig. 37) introduced from ‘la rade d’Amboine ’ the new as he also illustrated (pl. 5 fig. 1) a Hyalea ‘tricuspidata ’, where ob - (gymnosome) species Clio pyramidalis (not a homonym viously ‘ tridentata ’ is meant. In WoRMS (Gofas, 2014) this of Clio pyramidata Linné, 1767) . The name Clio pyrami - Bowdich citation is, erroneously and with the incorrect year dalis was later on discussed by Souleyet, in Rang & 1820, interpreted as Clio cuspidata (Bosc, 1801). Souleyet (1852 : 80, pl. 11 fig. 13) and also by Pruvot-Fol

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D’Orbigny (in de la Sagra, 1841 : 83) again described Cleodora pyramidata , mentioning it specifically also from the Caribbean (‘Antilles’) . In a footnote on p. 83 the specimen illustrated by La Martinière (1797) is in - troduced as a new species: ‘ L'espèce figurée par Lapey - rouse pl. 20, f. 1-3, que nous avons rencontrée dans le grand Océan , nous paraît distincte de celle de l'océan At - lantique, et nous la nommons Cleodora Lamartinieri ’. This name is repeated on one of the last pages of ‘Voyage dans l’Amérique ’ (1846: 688), as the caption of plate 8 figs 30-31, which, however, represent a speci - men with strongly concave carinae, very much differ - ent from La Martinière’s illustrations (see Figure 6 above). This discrepancy led van der Spoel (1967) to citing ‘d’Orbigny (184 1, non 184 6)’ for Clio pyramidata f. pyramidata , but ‘d’Orbigny (184 6, non 184 1)’ for C. pyramidata f. lanceolata . Fig. 11. De Blainville’s 1827 illustrations of Cleodora brownii . Two fossil pteropod species were introduced by Calandrelli (1844) from the Pliocene of Monte Vati - cano, Roma, Italy, namely Cleodora vaticana and C. ric - (1932 : 508), who both expressed some doubts about the cioli . The former, however, illustrated in Calandrelli’s validity of that species. Tesch (1913 : 130) did not give a upper figures A-B , is a subjective junior synonym of clear opinion on this taxon, but referred to it without Clio pyramidata , and represents the form with concave comments at the end of his text on the genus Clione . carinae , as explained in Janssen (1995 : 90 fig . 1 ). In According to van der Spoel (1976: 130) , however, Calandrelli’s illustration not only the two new species Quoy & Gaimard’s name Clio pyramidalis is a synonym were drawn, but also (his fig. C) a specimen of of the gymnosome species Clione limacina (Phipps, ‘Cleodora caudata with reference to Rang (1829, pl. 49 1774) or its f. antarctica (Smith, 1902). This seems fig . 1)’ . The reference is erroneous, it is not a copy of 89 doubtful, as according to van der Spoel’s (1976 fig . 221) Rang’s illustration, but it is based on Péron & Lesueur distribution map of that colder water species neither C. (1810, pl. 2 fig. 14, as ‘Cléodore pyramidale’), and re - limacina , nor its f. antarctica occurs in the tropics near drawn by Calandrelli. That drawing indeed resem - Amboina. See above for Bowdich’s (1822) and Zilch’s bles several earlier illustrations of C. pyramidata , as 1959 erroneous use of the name Clio pyramidalis . interpreted in those times. Calandrelli’s figures are D’Orbigny (1834 : 113; 1836, pl. 7 figs 25-32), in his copied herein as Fig . 12. ‘Voyage dans l‘Amérique méridionale ’, described the pres - ent species as ‘Hyalæa pyramidata Linn.’, interpreting the genus Hyalæa in a very wide sense ( 1834: 77), in - cluding a number of earlier names in it as synonyms, among which also Clio , Carolina [sic ], Creseis and Cleodora , but accepting several of these as a subgenus . For Hyalæa (Cleodora ) pyramidata a number of earlier references are given, among which also La Martinière’s 1797 illustrations. D’Orbigny distinguished for Hyalæa pyramidata the varieties A (‘ très élargie latéralement ’) and B (‘ presque pyramidale ’) and gave different geographical distributions, but mentioned his var. B (p. 11 5) exclu - sively from ‘le grand Océan austral ’. Of the earlier illus - trations of Browne (1756 ; which d’Orbigny considered to represent a juvenile specimen) and La Martinière Fig. 12. Cleodora vaticana Calandrelli, 1844 (A, B) and ‘ Cleodora (1797) d’Orbigny wrote ’ C’est évidemment la variété B ’. caudata ’(C), from Calandrelli (1844). Fig. A represents the exter - Although he gave (1834: 114) the reference to nal mould, Fig. B the internal one, both of the same specimen. Lesueur’s 1813 description of Hyalea lanceolata and Fig. C according to the author after Rang (1829), but in reality is recognised it as his ‘variété A ’ he did not use Lesueur’s redrawn after Péron & Lesueur (1810, pl. 3 fig. 14, as ‘Cleodore name for it, agreeing with his habit to indicate a ‘var - pyramidale’). iété’ with a single character only .

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Gould (185 2: 488; 1856, pl. 51 fig. 605, 605a-b) intro - According to its short description , Cleodora occidentalis duced Cleodora exacuta from the Pacific Ocean (44°N, Dall (1871 : 140), not illustrated by the author, is a form 154°W). We agree with van der Spoel (1967 : 69) that of Clio pyramidata with concave carinae. It was col - his large figure, here reproduced in Fig . 13, represents lected in the Pacific Ocean, off the California coast, at the form of Clio pyramidata with concave carinae . coordinates 33°N, 130°W. However, his fig. 605a, showing the transverse shape In Reeve’s Conchologia Iconica , G.B. Sowerby (1878, of the shell, does not really agree with C. pyramidata , Pteropoda pl. 4 fig . 24) described Cleodora lobata (Fig . 15 the transverse shape of which is much more triangu - herein) and compared it to d’Orbigny’s (1834-1836) lar. Hence, in our list of synonyms in the systematic Hyalæa pyramidata var . A, while considering it a sepa - chapter below we accept this name with a query. rate species. The rather poor illustration represents a Clio pyramidata with concave carinae. In the same work (fig. 25) also Cleodora pyramidata is illustrated, equally showing a specimen with concave carinae. His fig. 22 is named Cleodora australis , with reference to d’Orbigny (183 4, pl. 8 figs 12-14 ([incorrect, meant were figs 9-11] ). According to van der Spoel (1967 : 71 ), however, Sowerby’s specimen belongs to C. pyramidata forma sul - cata (Pfeffer, 1879), whereas the reference in d’Orbigny, again according to van der Spoel (1967 : 72), represents C. pyramidata forma antarctica Dall, 1908.

Fig. 13. Clio exacuta Gould, 1851 from the North Pacific Ocean. Size indicated as ‘length about half an inch, breadth equal to about two thirds the length’.

90 Souleyet, in Rang & Souleyet (1852 : 50, pl. 5 figs 7-9), described Cleodora pyramidata , but considered Péron & Lesueur (1810) to be the authors of this combination although referring to Browne’s (1756) and Linné’s (1767) descriptions in the synonyms. He also consid - Fig. 15. Cleodora lobata G.B. Sowerby, 1878, from the Atlantic ered Cleodora lanceolata Lesueur to be a synonym , as in Ocean, dimensions not indicated. his opinion (obviously based on d’Orbigny, 1834 : 114) it is just a more adult form of C. pyramidata . The illus - trations (copied herein as Fig . 14) represent a well-de - Boas (1886 : 69, 203) , discussing ‘ Cleodora pyramidata ’, veloped specimen of the shell form with concave introduced three named varieties: var. angusta (agree - carinae and, contrary to their text, indicated in the ing with d’Orbigny’s var. B), var. lata (d’Orbigny’s var. plate caption as Cleodora lanceolata . The genus Clio was A) and var. convexa . He also gave differing geographi - discussed on p. 77ff, but included only the gymno - cal areas for these forms. The ‘variety convexa ’ for a some species. long time was indeed considered to be a form of Clio pyramidata (see van der Spoel, 1967, 1969) , but is currently accepted as an independent species (Bé & Gilmer, 1977; Richter, 1979 ; Rampal, 2002; Janssen, 20 07 a, b) not occurring in the Atlantic Ocean. In his list of synonyms Boas referred to ‘Hyalæa lanceolata Lesueur ’, but curiously did not use it as an older name for his variety lata . On plate 6 figs 96a-g a series of seven specimens is represented in outline (Figure 16 herein ), showing an increasing degree of concavity of the carinae. Although there is not an Fig. 14. Cleodora lanceolata Lesueur, after Souleyet, in Rang & obvious boundary between them Boas considered Souleyet, 1852. numbers 96a-f to belong to his variety angusta and only 96g as variety lata.

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Fig. 16. Cleodora pyramidata after Boas (1886, pl. 6 fig 96) figs a-f = var. angusta , and fig. g = var. lata .

From the late Miocene Navidad Formation of Chili Mediterranean and the West Pacific Ocean. Clio promaucana Philippi (1887 : 112, pl. 13 fig. 18) was Tesch’s diagnosis of subspecies Clio pyramidata con - described. Judging from the poor illustration (herein vexa is erroneous. The clearly distinguishing character - copied as Fig. 17) we interpret it as a probable junior istic of the double-lined carinae is not mentioned and synonym of Clio pyramidata . According to Nielsen also his distributional data do not agree with the real (2003 : 198) the type specimen is lost. area of occurrence of Clio convexa. Based on an unpub - lished opinion of McGowan (1960), C. convexa was raised to full species level by Bé & Gilmer (1977: 773). A fossil species from the Pliocene Bowden Beds of Jamaica was introduced as Cleodora bowdenensis Collins (1934 : 201, pl. 12 figs 3-7 ). It was synonymised with Clio pyramidata by Robba (1977 : 600) and with C. pyramidata f. lanceolata by Janssen (1999 : 21, pl. 3 figs 13-17). The holotype is re-illustrated herein as Fig. 18. 91 Two drawings of the form with concave carinae were published by Korobkov (1955: 430 fig. 195) repres - ening an Italian Pliocene specimen (a) and a recent one (b), indicated with the name Clio pyramidata (Fig. 1 9). Fig. 17. Clio promaucana Philippi, 1887, holotype, shell height 6.5 mm (after Philippi, 1887).

Twentieth century and present -day concept

Tesch (1913) distinguished Clio pyramidata with three subspecies. For Clio pyramidata pyramidata , a long and rather uncritical list of synonyms is given, among which Browne (1756), Linné (1767, with C. caudata and ‘? C. retusa ’ included ), and finally also Boas’ var. angusta . The latter form is illustrated in Tesch’s fig. 31A-C, of which fig. 31A is a reproduction of Boas’ fig. 96a. The author repeated Boas’ distribution data saying that this form is restricted to the North At - lantic between 40 °N and 60°N and eastern parts of the South Pacific, and adding ‘occasionally to Davis Strait and Spitzbergen ’. Tesch’s second subspecies is Clio pyramidata lanceo - lata , for which also several synonyms are given, start - ing with Lesueur (1813) and concluded with Boas’ Fig. 18. Holotype of Clio bowdenensis (Collins, 1934). Upper row: (1886) var. lata . As distribution is given the warmer after Collins (1934), lower row: after Janssen (1999). parts of oceans between 40°N and 30°S, also in the

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The interpretation of two subspecies in the Atlantic Ocean, a northern one with virtually straight (forma ‘pyramidata’ ) and a southern one with strongly con - cave carinae (forma ' lanceolata '), is maintained in most of the recent literature (Rampal, 1975, 2002 , 2017 ; Janssen, 2007b, 2012). Several further taxa are distin - guished in the subantarctic region (Fig . 20).

Discussion Fig. 19. Clio pyramidata L., according to Korobkov (1955). Following the historical overview of Clio pyramidata ,it is clear that the current concept of ‘typical’ C. pyrami - The nomenclature as maintained by Tesch (1913) is not data being the form with straight shell sides finds its followed by van der Spoel (1967 : 67) who interpreted origin in the older literature and illustrations therein, Tesch’s three subspecies of Clio pyramidata as formae, but is here considered to be erroneous. In this respect together with several further, earlier described taxa. For especially the illustration in Browne (1756), here the North Atlantic, van der Spoel followed the interpre - copied in Fig . 2, is noteworthy, as it is exactly that fig - tation of Tesch as far as shell characteristics and distri - ure on which the taxon Clio pyramidata was based by bution are concerned. According to van der Spoel’s fig. Linné, 1767 (as it was the only reference given ). 348 Clio pyramidata f. pyramidata is distributed in the Browne’s illustration indeed seems to represent a North Atlantic between 4 5° and 6 5°N, whereas Clio specimen with straight shell sides. D’Orbigny’s (1834 : pyramidata f. lanceolata occurs between 50°N and 45°S 113 ) remark that Browne’s drawing represents a juve - worldwide, inclusive of the the Mediterranean, but ex - nile specimen is incorrect, as Browne himself, in his cluding the larger part of the Caribbean. Van der text on p. 386 mentioned the size of the species as ‘sel - Spoel’s (1967) interpretation of Clio pyramidata f. convexa dom exceeds half an inch’, which certainly does not follows Tesch’s morphological data. pertain to very juvenile specimens .

92

Fig. 20. Van der Spoel’s (1967 fig. 348) distribution map of Clio pyramidata formae.

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In our opinion Browne’s illustration is incorrect and N s : I e * * r 1 1 2 1 1 1 1 1 5

too much schematised . A form of Clio pyramidata with O 2 1 u t D c t such straight carinae does not occur in the area from i a P y

where Browne’s illustrated specimen was collected, r o t 7 8 9 0 1 2 3 4 5 3 9 i 4 4 4 5 5 5 5 5 5 2 s which is the Caribbean Sea near the island of Jamaica. 8 6 6 6 6 6 6 6 6 6 9 7 o 1 1 1 1 1 1 1 1 1 1 8 2 p 0 0 0 0 0 0 0 0 0 4 O 9 e

The typical form of C. pyramidata occurring in that 1 1 1 1 1 1 1 1 1 0 r 2 C F l P H H H H H H H H H a K t

area has shells with distinctly concave sides, de - M M M M M M M M M M i g scribed in 1813 by Lesueur as Hyalea lanceolata , that i d d

we consider a subjective junior synonym of C. pyrami - a y r 1 3 1 2 1 2 2 1 2 1 2 0 0 0 0 0 0 0 0 0 0 0 D data. In the systematic chapter below we illustrate 11 e 8 8 6 6 6 8 6 1 0 6 0 d e 2 2 2 2 0 2 6 7 1 1 1 o h 4 A A A A A A t U U c A

specimens from Jamaica that invariably demonstrate M t A r G G n M M M M M M A M i y o y y y y g g g g e these clearly concave carinae of their shell (Figs 24 & u p e r p p h c a a a a p p l L L L L S L L L L L L L b C a C C C C C C C C 25) . We examined the relationship of the two shell C C l i a

forms of C. pyramidata and conclude that the form v a e

with concave sides (occurring in the warm waters of r 2 a 1 2 2 0 0 1 1 2 2 1 3 0 0 1 k _ _ 0 0 0 0 0 0 _ _ s 0

the Atlantic, Caribbean and Mediterranean) should be 0 i 1 ______0 6 _ 0 r 7 6 6 6 8 8 8 1 6 6 1 e _ 0 2 2 2 2 2 _ _ t 1 e considered typical ' C. pyramidat a' and the form with _ 1 ______s 4 8 _ 1 d 0 a 2 1 R R R R R R 0 R o 1 T 1 T c n A A A A 1

straight sides could be indicated as ' angusta ' Boas, A A A 1 a g M M U M M M M M M U M n h _ _ _ _ A _ _ A G t 1886 , but at subspecies rank only. _ o G r r i _ g g g g _ c _ _ r L s y y r e n n n n r w y r y u p p a a a a y _ c p _ _ d _ _ _ _ p l l l p l l l l _ _ e _ l l _ t l C C C C C C C Molecular methods l a C C C c C i d n i n s o 0

To test whether forms ' pyramidata ' and 'angusta ' are r i 1 6 6 8 8 8 0 6 6 t 0 6 e 7 2 2 2 2 2 1 6 1 a 1 b genetically different , genomic DNA was extracted t S m u

from soft tissue of nine Clio pyramidata specimens n e O O O O O O O r 1 1 4 8 e u using the DNA barcoding pipeline at Naturalis Biodi - C C C C C C C 0 0 2 1 t s E E E E E E E 1 1 i c T T ------i 1 1 u R R R R R R R

versity Center (described in detail in Wall-Palmer et P r M M U U . A A A A A A A C I A A G G

al. , 2018). A fragment of ~600 basepairs (bp) of the mi - e 93 M M M M M M M s a

tochondrial Cytochrome Oxydase I (COI) gene was d i - i W W W W W W W W W W W x ' ' ' ' ' ' ' ' ' ' ' e g 5 7 7 0 0 9 9 9 2 0 7 O amplified using the primers from Geller et al. (2013) d n 4 2 0 2 2 2 2 2 4 0 1 u e ° ° ° ° ° ° ° ° ° ° ° o t 6 5 1 9 9 7 7 7 8 5 3 L m

according to methods described in Wall-Palmer et al. 8 8 3 2 2 2 2 2 2 2 3 o r

(2018). Nucleotide sequences of COI were examined h e c S N N N N N N N N N N o ' d ' ' ' ' ' ' ' ' ' ' t 0 u 0 0 0 4 4 5 5 5 9 4

and aligned using the MUSCLE algoritm in MEGA 6 y t 5 1 3 1 0 0 3 3 3 4 3 i ° ° ° ° ° ° ° ° ° ° ° C t 2 6 9 7 8 8 2 2 2 7 1 a 1 n

(Tamura et al., 2013). A Maximum Likelihood (ML) 1 1 5 4 4 4 4 4 1 5 L o

analysis was used to estimate evolutionary relation - d e e s t a a 8 3 2 5 5 7 7 7 8 7 9

ships among taxa (Felsenstein, 1981) in RaxmlGUI 1.3 b d 1 2 1 2 2 2 2 2 0 2 1 ------s i n 4 4 6 6 6 6 6 6 0 0 6 s

(Stamatakis, 2006; Silvestro and Michalak, 2012). For o 0 0 0 0 0 0 0 0 1 1 0 i y ------t l 1 1 4 4 4 4 4 4 4 8 4 c a 1 1 0 0 0 0 0 0 1 0 0

this the GTR (General Time Reversible) model was se - e n 0 0 0 0 0 0 0 0 0 0 0 l a l 2 2 2 2 2 2 2 2 2 2 2 o c

lected, using codon-specific evolutionary rates. i C t e

Five specimens morphologically resembled ‘pyra - n e 5 7 g 1 1 o

midata ’ with concave shell sides and four specimens re - l 0 0 y 2 2 , , h y y y y y y y y y e . . l l angusta c

sembled ‘ ’ with more straight shell sides, and p d d d d d d d d d a a a n u u u u u u u u u e a t t t t t t t t t t t e d r s s s s s s s s s e e

were numbered according to the degree of concavity n i i e s s s s s s s s s e e o f i i i i i i i i i i d g g e h h h h h h h h h n of shell sides (Fig . 21 ). Specimens from the Caribbean e d d i R T T T T T T T T T s i i l r r u o Sea were collected during the GU1101 expedition in r r s v u u e a B B c C

2011 ( N=2 ' pyramidata ' from 2 stations). Specimens from n , e a u t a the North Atlantic Ocean were collected from three lo - q a a a a a e t t t t t m s a a a a a a a a a s t t t t o d d d d d o

cations during the MAR-ECO cruise in 2004 ( N=4 ' an - s s s s e i i i i i s i i l u u u u o c m m m m m g g g g C c e a a a a a a .

gusta ' and N=2 ' pyramidata '), and from one location n n n n e f t r r r r r a p a a a a 6 a y y y y y v o h S 0 t r d p p p p p a a a a i 1 t t t t u further south during the Atlantic Meridional Transect u w a a a a a p a a a a & c p t t t t t E e s e d d d d i b a a a a a i i i i , u r y . v d d d d d c 24 (AMT24) in 2014 (N=1 ' pyramidata '; Table 1 , Fig . 21 . a i i i i i d m m m m r o t m i o a a a a e a l a i m m m m m o r r r r l y v a a a a a C y y y y n

Outgroup taxa for phylogenetic analyses were Clio cus - r m r r r r r C O p p p p o o d y y y y y . / x s p p p p p o o o o i i i i 1 1 a

pidata (Bosc, 180 1) (N=1) and Clio recurva (Children, o l l l l o o o o o 6 c i i i i i T e C C C C l l l l l 0 l e 5 C C C C C b h 1823) (N=1; Table 1; Burridge et al., 2015, 2017 ). All lo - . a 0 T 1 cality data are specified in Table 1 . T

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Fig. 21. Overview of the nine Clio pyramidata specimens used for molecular phylogenetic analysis. Specimens are arranged based on de - gree of concavity of shell sides. Bar size is 5 mm.

Molecular results Conclusions

Based on the phylogenetic analysis of CO1 gene se - We conclude that the interpretation of ‘typical’ Clio quences, support was found for monophyly of all nine pyramidata has been erroneous throughout history, C. pyramidata specimens (Fig . 23). The shell forms and that the taxon Hyalea lanceolata , introduced from 'pyramidata ' and 'angusta ' could not be distinguished, the Mediterranean by Lesueur (1813), is a subjective nor was there any support for a distinction between junior synonym of Clio pyramidata Linné, 1767. We sta - specimens from the Caribbean Sea and specimens bilise nomenclature by the designation of a neotype from the open North Atlantic Ocean. It is noteworthy for Clio pyramidata on the basis of a specimen from the that our sampling included intermediate shell forms type locality, the sea around Jamaica (see sytematic (Fig. 21) and that both shell forms were found in the part below ), ‘ with the express purpose of clarifying its same zooplankton sample (station 28 from the taxonomic status ’(ICZN art. 75.3.1). MARECO expedition, Fig. 22). Hence, notwithstand - It is evident that the form occurring in the northern 94 ing our limited sampling of specimens and molecular Atlantic Ocean can no longer be indicated as Clio pyra - markers, we suggest that different shell forms do not midata (f.) pyramidata as it distinctly differs from neo - represent distinct species typical C. pyramidata by its straight rather than concave sides of the shell. For this morphotype the name angusta Boas,1886 , is available and to avoid any future misunderstanding we designate a lectotype from Boas’ syntypical material housed at the Natural History Museum of Denmark, Copenhagen (see sys - tematic part, below) . The even older name Clio lamartinieri d’Orbigny, 1841, based on La Martinière’s 1797 illustration, repre - sents a species from the NE Pacific Ocean, where a form of Clio pyramidata with straight carinae does not occur. Therefore, and also because of d’Orbigny’s 1841 and 1846 differing interpretations, we consider the name C. lamartinieri a nomen dubium . Also we cannot exclude the possibility that it, in fact , represents Clio convexa (Boas, 1886). Our molecular results demonstrate that the shell forms ‘pyramidata ’ and ‘ angusta ’ are unlikely to repre - sent distinct species, meaning that the latter name can only be applied at subspecific rank.

Fig. 22. Overview of sampling locations of Clio pyramidata speci - mens (N=9) used for molecular phylogenetic analysis. White square = ' angusta '; Black square = ' pyramidata '; Half-white square = ' angusta ' and ' pyramidata ' at the same location.

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Fig. 23. Phylogeny based on mitochondrial Cytochrome Oxidase I sequences of Clio pyramidata (N=9) and outgroup taxa C. cuspidata (N=1) and C. recurva (N=1). Numbers 1-9 follow numbers in Figure 22.

Systematics ramidale’ (= Clio pyramidata Linné, 1767). ? Balantium Children , 1823: 220. Monotype Balantium recurvum Order Pteropoda Cuvier, 1804 Children , 1823. 95 Suborder Euthecosomata Meisenheimer, 1905 ? Balantium (Flabellulum) Bellardi , 1873: 32 . Type species Balan - Family Cliidae Jeffreys, 1869 tium braidense Bellardi, 1873 (here designated). ? Balantium (Poculina) Bellardi , 1873: 33. Homonym of Poculina Cleodoridae Gray, 1840 : 148, 155 ( designated nomen oblitum by Gray, 186 8: 742 (Gastropoda, Calyptraeidae). Bouchet & Rocroi , 2005: 51). Euclio Bonnevie , 1913: 20. Objective junior synonym of Clio . Cliidae Jeffreys , 1869: 118; ICZN, 2006, Opinion 2133. ? Proclio Hubendick , 1951: 3. Monotype Proclio subteres Huben - Clionae van der Spoel , 1967: 31, 57 ( partim ). dick (1951: 6 fig. on p. 7, pl. 1 figs 1, 2) = Clio antarctica (Dall, Clioidae Bouchet & Rocroi , 2005: 51. 1908). Cavoliniidae Bouchet et al . 2017: 357 ( partim ). ? Bellardiclio Janssen , 2004: 114 . Type species by original desig - Non Clioidae Menke , 1828: 5 (as Cliodinae), currently Clionidae nation Hyalæa cuspidata Bosc, 180 1. (Gymnosomata). Clio pyramidata Linné, 1767 s. str . Note – Cliidae was synonymised with Cavoliniidae (Figs 24, 25) by Bouchet et al . (2017: 357), based on molecular work of Burridge et al . ( 2017). The general morphol - Below we list the various names and name combina - ogy of species earlier included in Cliidae, however, is tions used for the typical form of this species during so analogous and different from other time. For more extensive synonymy see van der Spoel that we suppose that further detailed and integrative (1967) and Janssen (2012). taxonomic research eventually will result in a more robust subdivision validating this family. Clio vagina triquetrâ pyramidata, ore oblique truncato . The smaller Clio , with a trilateral sheath – Browne , 1756: 386, pl. 43 fig . Clio Linné, 1767 1. [non binom.] Clio pyramidata Linné , 1767: 1094. Clio Browne , 1756: 386 ( unavailable, pre-Linnean). ? Clio caudata Linné , 1767: 1094. Clio Linné , 1767: 1094. Type species Clio pyramidata Linné, 1767 ? Clio retusa Linné , 1767: 1094. by subsequent designation of Gray (1847 : 203). ‘Insecte’ – La Martinière, 1787a : 207, pl. 11 figs 1-3 . Cleodora Péron & Lesueur , 1810: 66. Monotype ‘le Cléodore py - Hyalæa pyramidata – Bosc , 1801: 241, pl. 9 figs 1-3; Lamarck,

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1819: 290. bryonic shell is more elongated than in the forma pyra - ‘Cléodore Pyramidale ’ – Péron & Lesueur, 1810 : pl. 2 fig .. 14. midata [= angusta herein] . The radula (formula 1-1-1) Hyalea lanceolata Lesueur , 1813: 284, pl. 5 fig . 3. shows about 6 transversal rows. Тhе shell length is Cleodora pyramidalis [sic ] – Bowdich , 1822: 23, 56, pl. 5 fig . 2. about 20 mm ; the maximal width about 15 mm .” ? Cleodora Brownii de Blainville , 1824: 272; 1825: 481; 1827, pl. Material examined . — Apart from the neotype 46bis figs 1, 1a-b). specimen, 1 0 additional specimens from the neotype Hyalæa (Cleodora) pyramidata Linn. Var. A – d’Orbigny , 1834: 113; locality, RGM 777 445 (Fig . 25) (see caption of Fig. 24 1836, pl. 7 figs 25-32. for details ). ? Cleodora Lamartinieri d’Orbigny , 1841: 83 (footnote). Discussion . — The lot of 1 1 specimens of Clio pyra - Cleodora Vaticana Calandrelli , 1844: 13 figs A, B. midata Linné, 1767 from the neotype locality Jamaica ? Cleodora exacuta Gould , 1852: 488; pl. 51 fig, 605, 605a-b (1856). (Figures 24-25) demonstrates convincingly that speci - Clio occidentalis Dall , 1871: 140. mens of C. pyramidata from that area have strongly Cleodora pyramidata (L.) var. lata Boas , 1886: 72, 203, pl. 9 figs 96g. concave shell sides. Therefore the form of this species ? Clio promaucana Philippi , 1887: 112, pl. 13 fig . 18. with straight or almost straight sides, occurring in the Clio (Clio) pyramidata Linné – Pelseneer , 1888: 63. northern Atlantic Ocean, indicated as C. pyramidata (f .) Clio (Euclio) pyramidata Linné var. lata – Bonnevie , 1913: 23 fig. pyramidata by several authors (e.g., Tesch, 1913; van 22B. der Spoel, 1967; Janssen, 2012) cannot be considered to Clio pyramidata lanceolata (Lesueur) – Tesch , 1913: 36 fig. 32A-C. be typical C. pyramidata . The form with strongly con - Cleodora bowdenensis Collins , 1934: 201, pl. 12 figs 3-7. cave carinae currently is usually indicated in literature Euclio pyramidata (Linné) form lata – Tesch , 1946: 14, pl. 2 figs with the name C. pyramiata (f.) lanceolata (Lesueur, 11a-f. 1813) which is herewith demonstrated to be a subjec - Euclio pyramidata lanceolata (Lesueur), 1813 – Tokioka , 1955: 62, tive junior synonym of C. pyramidata Linné, 1767. As pl. 8 figs 11-13. this is one of the most common and widely distrib - Clio pyramidata Linnaeus, 1767 forma lanceolata (Lesueur, 1813) – uted pteropod species , a correct naming of the species van der Spoel , 1967: 68 figs 50-54, 60 (with extensive syno - is of importance. nymy). Clio (Clio) pyramidata Linnaeus, 1767 forma lanceolata (Lesueur, Clio pyramidata angusta (Boas, 1886) 96 1813) – Janssen , 2012: 44 figs 16A-D, 17A-F, 49J-L (with addi - (Fig. 26) tional synonymy) . Cleodora pyramidata (L.) var. angusta Boas , 1886: 72, 203, pl. 6 fig. Neotype – RGM 777 444 figure 24 a-d, here designated. 96a-f. Neotype locality – Jamaica, East Channel, bottom Clio (Euclio) pyramidata Linné (var.) angusta – Bonnevie , 1913: 23 sample no. Est 875F, leg. Ivan Goodbody, R/V East - fig. 22A. ward, 17 February 1972; don. M.K. Webber, November Clio pyramidata pyramidata L. – Tesch , 1913: 35 fig. 31A ( non 2016. Linné) . Description (after van der Spoel, 1967: 69 -70, as Clio pyramidata Linnaeus, 1767 forma pyramidata Linnaeus, 1767 – Clio pyramidata forma lanceolata ). — “Shell nearly van der Spoel , 1967: 67 figs 48a-b, 49 ( non Linnaeus ). straight, pyramidally shaped, thin and transparent. Clio pyramidata angusta – Bé & Gilmer , 1977: 766. Тhe lateral ribs are thickened, and very strongly di - verging. Тhe dorsal rib is straight and protrudes dis - Description (after van der Spoel, 1967: 6 8, as Clio pyra - tinctly above the shell aperture. Тhe lateral ribs are midata forma pyramidata ). — “Shell, hyaline sometimes very long too, but never end in free processes. Тhe with а reddish shine and thin, straight pyramidally cross-section through the shell is rounded near the shaped. Th е two lateral ribs are thickened and only posterior top, triangular in the middle and the ante - slightly diverging and bent. Transverse striation and rior half of the shell. Anteriorly the ventral surface is growth lines are distinct. Cross-section triangular in hollow so the cross-section assumes а lunar shape.' all parts except the most posterior part. Тhe dorsal rib Тhis hollow shape of the ventral side is effected b у the protrudes slightly аbov е the shell aperture. Тhe latero- lateral ribs bent ventrally in the upper рart . At both dorsal sides are both provided with three swellings оf latero-dorsal sides of the shell three longitudinal ribs the shell, and one longitudinal rib runs over the ven - are present and оnе rib is found оn the ventral side. tral shell surface. Тhe greatest width оf the shell, the Тhe transverse striation and growth lines are distinct aperture width, is found аbov е the middle of the shell. but usually not as pronounced as in the f оrm а pyrami - Тhe lateral ribs оf the shell are rounded, especially data [= angusta herein] .Тhе greatest width of the sh еll near the embryonic shell. Тhе embryonic shell is is found near the middle of the shell. Тhe lateral ribs droplet-shaped and provided with а small cusp. Тhe are rounded пеаг the posterior shell top. Тhе oval em - radula (formula 1-1-1) is composed оf about 7 rows.

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Fig. 24. Clio pyramidata Linné, 1767. Neotype, RGM 777 444. Jamaica, East Channel, boBom sample nr Est 875F, leg. Ivan Goodbody, R/V Eastward, 17 February 1972; don. M.K. Webber, November 2016; a: apertural, b: dorsal, c: right lateral, and d: ventral views.

97 Тhe shell length is about 20 mm; its width about 10 NHMD 199251 - 58°27'N 26°43'W (2 specimens, one of which il - mm. ” lustrated in Fig, 2 6b). Lectotype (here designated) .—NHMD 199348, NHMD 199261 - 59°3'N 13°32'W; 14 specimens . North Atlantic Ocean, 58°N 28°W (Figure 2 6a). NHMD 199331 - 58°27'N 26°43'W; 18 specimens, one of which The following syntypical lots, now paralectotypes, illustrated in Fig. 2 6c. are available: NHMD 199348 - 58°N 28°W (lectotype locality) ; 17 specimens , two of which illustrated in Figs 26a and 26d).

Fig. 25. Clio pyramidata Linné, 1767 s. str., additional specimens (all dorsal views) from the neotype locality (see caption of Fig. 24 for de - tails), RGM 777 445. The protoconch is only preserved in specimen e.

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NHMD 199365 - 58°26'N 19°WE; 23 specimens, includes many sequencing. Annelies Pierrot-Bults (guest researcher at juveniles. Naturalis Biodiversity Center, Leiden, NL) provided NHMD 1993375 - 59°17'N 8°W; 6 specimens . specimens from the MAR-ECO expedition. Tom NHMD 199380 - 59°N 18°W; 28 specimens . Schiøtte [Natural History Museum of Denmark (Zool - NHMD 199399 - 60°18'N 16°48'W; 2 specimens . ogy), Copenhagen] made syntype material of Boas (1886) available to us. Steve Tracey (Natural History Discussion . — Clio pyramidata angusta , as far as cur - Museum, London, U.K.) provided copies of Lesueur’s rently known, is restricted to the northern Atlantic (1813) drawings. Edmund Gittenberger (Naturalis Bio - Ocean, north of c. 45 °N and may have developed from diversity Center, Leiden, NL) shared his views on the C. pyramidata s. str. under the influence of lower sea - application of the concepts of species and subspecies. water temperatures. Locally these two shell forms Eduard F. de Vogel, of the same institute, made the may occur sympatrically and intermediate shell forms stacking photography images of figures 24-26. Jaap occur. Furthermore, so far no genetic differences are van der Voort (Venne, Germany) assisted in the litera - found between the two shell forms and thus they are ture searches. We thank Deborah Wall-Palmer for her unlikely to be reproductively isolated. careful reading of the manuscript and many useful Several authors (e.g., d’Orbigny, Boas, Tesch) men - comments. tioned similar occurrences with more or less straight References carinae also from the Pacific Ocean and illustrations of similar specimens can be found in various websites. Adams, H. & Adams, A. , 1853-1858. The genera of recent Mol - However, it cannot be excluded that such specimens lusca; arranged according to their organization. John van from the Pacific or Indian Ocean are in fact Clio con - Voorst, London: 1: 1-256, pls. 1-32 (1853); 257-484 pls. 33-60 vexa (Boas, 1886 ), a species at first glance very similar (1854); 2: 1-92, pls. 61-72 (1854); 93-284, pls. 73-96 (1855); 285- in shape, but easily distinguished by the double-lined 412, pls. 97-112 (1856); 413-540, pls. 113-128 (1857) , 541-660, carinae with a square transverse section and a differ - pls. 129-138 (1858). ent shape of its larval shell. Bellardi, L. , 1873. I molluschi dei terreni terziari del Piemonte e della Liguria, 1. Cephalopoda, Pteropoda, Heteropoda, Gast - Acknowledgements ropoda (Murcidae et Tritonidae). — Memorie della Reale 98 Accademia delle Scienze di Torino (2) 27: 1-264, 15 pls. We are grateful to Mona K. Webber (University of the Blainville, [H.M.D.] de , 1817. Cléodore (Malacoz.), Cleodora . — West Indies, Discovery Bay Marine Laboratory , Dis - Dictionnaire des Sciences Naturelles 9: 386-388. covery Bay, Jamaica) for providing the neotype lot . Blainville, H.M.D. de , 1821. Mémoire sur le genre Hyale. — Rebeca Gasca (El Colegio de la Frontera Sur, Chetu - Journal de Physique, de Chimie, d'Histoire Naturelle et des mal, Mexico) provided fresh Caribbean specimens for Arts 93: 81-97.

Fig. 26. Clio pyramidata angusta (Boas, 1886); a: lectotype; b-d: paralectotypes; c-d: transitional forms to Clio pyramidata s. str. Northern At - lantic Ocean, a and d from NHMD 199348, 58°N 28°W, b from NHMD 199251, 58°27'N 26°43'W, c from NHMD 199331, 58°27'N 26°43'W.

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Blainville, [H.M.D.] de , 1824. Mollusques, Mollusca. (Mala - Todd, J.A., Young, J.R., Osborn, K.J., Menken, S.B.J., & Peij - coz.). — Dictionnaire des Sciences Naturelles 32: 1-392 nenburg, K.T.C.A. , 2017. Time-calibrated molecular phy - Blainville, H.M. Ducrotay de , 1825-1827. Manuel de malacolo - logeny of pteropods. — PloS ONE 12 (6): e0177325. gie et de conchyliologie; contenant: 1e Une histoire abrégée https://doi.org/10.1371/ journal.pone.0177325 de cette partie de la zoologie; des considérations générales Calandrelli, A. , 1844. Scoperta di due nuove specie di conchi - sur l'anatomie, la physiologie et l'histoire naturelle des ma - glie fossili fatta dal sig. Giuseppe Riccioli nell'argilla del Va - lacozoaires, avec un catalogue des principaux auteurs qui ticano. Lettera diretta al chiarissimo sig. 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